GeolCarp_Vol65_No5_375

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, OCTOBER 2014, 65, 5, 375—386 doi: 10.2478/geoca-2014-0026

Introduction

Post-sedimentional erosional processes and Cretaceous sedi-
mentation removed or buried the Upper Jurassic deposits of
the  Bohemian  Massif.  The  depositional  conditions,  ecosys-
tem composition, and biogeographical affinity of this region
thus remain weakly known. However, a few outcrops captur-
ing  the  Upper  Jurassic  deposits  are  still  exposed  or  fossil
specimens from older outcrops are preserved in museum col-
lections. Such findings allow us to fill the gap in distribution
patterns  of  Upper  Jurassic  marine  organisms.  Although  the
first Upper Jurassic ammonites were found in the Bohemian
Massif  in  the  1860s  (Lenz  1870;  Bruder  1881,  1882,  1885,
1886), the regional composition and temporal distribution of
ammonite  assemblages  as  well  as  their  compositional  rela-
tionships with adjacent geographical regions were poorly ex-
plored.  In  this  paper,  we  focus  on  systematic  revision  of
museum  collections  of  two  ammonite  families  of  high  bio-
stratigraphic  and  paleobiogeographic  significance  (Aula-
costephanidae and Cardioceratidae), and compare them with
co-eval  Tethyan  and  Boreal  ammonite  assemblages  from
other regions. We note that the majority of specimens in the
ammonite assemblages in the Bohemian Massif, represented
by Perisphinctidae, Ataxioceratidae, and Oppeliidae, will be
described in a separate study.

The systematics and paleobiogeographic significance of

Sub-Boreal and Boreal ammonites (Aulacostephanidae

and Cardioceratidae) from the Upper Jurassic of the

Bohemian Massif

JAN HRBEK

Institute of Geology and Paleontology, Faculty of Science, Charles University in Prague, Albertov 6, 128 43 Prague 2, Czech Republic;

hrbek.honza@seznam.cz

(Manuscript received March 1, 2014; accepted in revised form October 7, 2014)

Abstract: Upper Jurassic marine deposits are either rarely preserved due to erosion or buried under younger sediments
in the Bohemian Massif. However, fossil assemblages from a few successions exposed in northern Bohemia and Saxony
and preserved in museum collections document the regional composition of macro-invertebrate assemblages and thus
provide unique insights into broad-scale distribution and migration pathways of ammonites during the Late Jurassic. In
this paper, we focus on the systematic revision of ammonites from the Upper Oxfordian and Lower Kimmeridgian
deposits of northern Bohemia and Saxony. The ammonites belong to two families (Aulacostephanidae and Cardioceratidae)
of high paleobiogeographic and stratigraphic significance. Six genera belong to the family Aulacostephanidae (Prorasenia,
Rasenia, Eurasenia, Rasenioides, Aulacostephanus, Aulacostephanoides) and one genus belongs to the family
Cardioceratidae (Amoeboceras). They show that the Upper Jurassic deposits of the northern Bohemian Massif belong to
the Upper Oxfordian and Lower Kimmeridgian and paleobiogeographically correspond to the German-Polish ammonite
branch with the geographical extent from the Polish Jura Chain to the Swabian and Franconian Alb. Therefore, the
occurrences of ammonites described here imply that migration pathway connecting the Polish Jura Chain with habitats
in southern Germany was located during the Late Oxfordian and Early Kimmeridgian in the Bohemian Massif.

Key words: Oxfordian, Kimmeridgian, Czech Republic, biostratigraphy, paleobiogeography, ammonites,
Aulacostephanidae, Cardioceratidae.

Fig. 1. Schematic map with surface outcrops of Jurassic rocks and
Lusatian fault course in northern Bohemia (modified according to
Kopecký et al. 1963).

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In the Late Jurassic, the Sub-Mediterranean Realm was lo-

cated along the northern margin of Tethys (namely the Euro-
pean part in the vicinity of the Mediterranean Sea – southern
Spain,  southern  France,  peri-Carpathian  area  and  Balkan
area  in  the  east).  The  Sub-Boreal  Realm  ranged  from  Scot-
land, southern England, northern France, northern Germany,
northern Poland up to the southern part of the Russian plat-
form and Caspian area. The Boreal Realm was located north
of  the  Sub-Boreal  Realm.  It  was  characterized  by  high-lati-
tude,  cold-water  faunas  occurring  in  the  Canadian  Arctic,
Alaska, Greenland, Spitsbergen, the Barents Sea, and Siberian
area (Callomon 2003).

Aulacostephanids are typical of the Sub-Boreal Realm and

are used for zonal and subzonal subdivisions of the Sub-Bo-
real Upper Oxfordian and Kimmeridgian (Birkelund & Cal-
lomon 1985; Krymholts et al. 1988; Wright 2010). However,
aulacostephanids  also  occur  in  the  Sub-Mediterranean  and
Boreal regions. In contrast, Cardioceratids are typical of the
Boreal Realm at high latitudes, and occur less frequently in
Sub-Boreal  and  Sub-Mediterranean  realms.  The  proposed
global  stratotype  for  the  Oxfordian/Kimmeridgian  (O/K)
boundary, located at Staffin Bay, Isle of Skye in Scotland, is
characterized  by  numerous  aulacostephanid  occurrences
below  and  above  this  boundary  (Matyja  et  al.  2006;
Wierzbowski et al. 2006). This boundary is characterized by
the  replacement  of  the  aulacostephanid  Ringsteadia  and
Microbiplices by  Pictonia  and Prorasenia,  which  coincides
with the first appearance of the Boreal species Amoeboceras
(Plasmatites)  (Wierzbowski  et  al.  2006;  Glowniak  et  al.
2010; Wierzbowski 2010).

Materials and methods

This study is based on specimens known from the collec-

tions of the National Museum, Prague (NM-N), the Institute
of  Geology  and  Paleontology,  Faculty  of  Science,  Charles
University, Prague (CIGP) and the State Museum of Geology
and  Paleontology,  Dresden  (SMGPD).  This  paper  includes
descriptions  of  26  specimens  belonging  to  Aulacoste-
phanidae  and  Cardioceratidae,  mostly  coming  from  the
Czech Republic, partly including specimens recorded in Ger-
many.  The  ammonites  described  here  were  collected  in  the
vicinity  of  Šluknov,  Krásná  Lípa,  Doubice,  Brtníky  and
Kyjov,  in  the  former  Šternberk  quarry  (northern  Bohemia)
and in Hohnstein, Saxony (Germany) (Fig. 1).

Ammonite preservation does not allow the determination of

morphological  characters  in  the  majority  of  specimens.  Su-
tures and other surface structures are not preserved, and deter-
mination  of  the  body  chamber  and  phragmocone  part  is
hampered  by  poor  preservation  of  internal  moulds.  Determi-
nation to a genus level is not possible in some specimens owing
to strong shell deformation. However, the morphological char-
acters  of  shells  are  sufficiently  preserved  in  some  specimens
and allow comparison with aulacostephanids from Europe.

The following abbreviations are used in the ‘Measure-

ments’ chapter:

D (max)

maximum diameter preserved

D (phg-ad)

diameter of fully grown individual

D (phg)

diameter of final whorl (incomplete conch)

Wh/D

ratio of height and diameter

Wt/D

ratio of thickness and diameter

U/D

ratio of umbilicus and diameter.

Geological settings

Jurassic outcrops in northern Bohemia are exposed only at

several places as relicts of the epi-continental sea. These out-
crops are associated with the Lusatian Fault that is responsi-
ble for overturning of successions and probably contributed
to tectonic alteration and dolomitization (Eliáš 1981). Dur-

Fig. 2. The original lithostratigraphic scheme with lithotypes col-
lected near Krásná Lípa town. Two massive limestone beds (Bimam-
matum and Platynota Zones) represent the most significant parts with
high density of fauna (modified according to Bruder 1882, 1886).

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ing the younger phase of the Saxonic tectonic event, the
re-activation of the Lusatian tectonic zone pulled the Cado-
mian, Paleozoic and Jurassic sediments over the Cretaceous
sediments.

Carboniferous and Permian sediments form the bedrock of

the  Middle  Jurassic  Brtníky  Formation  (Suk  et  al.  1984),
which  reaches  up  to  20  meters  in  thickness  and  is  formed
predominantly  by  sandy,  fossil-poor  clastics.  The  onset  of
sedimentation probably corresponds to the Callovian (Bruder
1881, 1882; Kopecký et al. 1963; Eliáš 1981; Chlupáč et al.
2002).  The  Doubice  Formation  overlies  sandstones  of  the
Brtníky Formation and exceeds 100 m in thickness (Kopecký
et al. 1963; Eliáš 1981; Suk et al. 1984). It is predominantly
formed by micritic limestones that were deposited in a deep-
shelf  marine  environment.  The  lithological  composition  is
variable,  formed  by  interbeds  of  marly  limestones,  cherts,
marls, and sandy rocks – see origin profile (Fig. 2). Carbon-
ates are weakly to strongly dolomitized (due to influence of
the Lusatian tectonic zone and the Tertiary volcanism). The
Doubice Formation contains diverse fossil assemblages with
ammonites, belemnites, brachiopods, bivalves, sponges, for-
aminifera, echinoderms, bryozoans, annelids and fish (Bruder

1881, 1882, 1885, 1886; Eliáš 1981). A relatively high abun-
dance of sponges (also documented in limestone blocks pre-
served in the creek below the “Peškova stráň” locality) can
imply that some assemblages correspond to sponge megafa-
cies. However, a sedimentological analysis of Upper Jurassic
facies from northern Bohemia is not possible at present due
to the lack of surface outcrops.

Near Doubice, outcrops expose overturned successions

that are located close to the Lusatian fault. Pešek’s Hillside
near  Kyjov  is  a  small-scale  oucrop  formed  by  sandy  lime-
stones,  marly  carbonates  and  micritic  limestones.  Sparitic
limestones  contain  recrystallized  bioclasts,  mainly  sponges
and  bivalves.  Most  bioclasts  occur  in  micritic  limestones
(sponge spicules, brachiopods). Matrix and bioclasts (mainly
bryozoans, sponges and bivalves) are characterized by exten-
sive  recrystallization.  Some  thin-sections  show  shear-like
structures,  possibly  reflecting  partial  redeposition.  Peloids
and pellets-like structures are present (Fig. 3).

The location of coastal edge of this epicontinental sea is

unknown; possibly extensive Jurassic sediments were eroded
during the Early Cretaceous and at the beginning of the Late
Cretaceous. However, the existence of a connection between

Fig. 3. Thin sections obtained from the Late Jurassic lithotypes collected from the Peškova stráň hillside near Kyjov. A, B, C – Micritic
limestones: A, B – Predominantly almost oval-shaped peloid-like objects occur very abundantly with recrystallized organic relicts (sponges,
brachiopods); C – Strongly recrystallized bioclasts in a homogeneous matrix. D – Calcareous-clayey siltstone. Sparitic siltstone with
fragments of quartz suggests the possible contribution of terrestrial material. Recrystallized faunal relicts (bivalves, gastropods, corals,
sponges) occur at a lower density in comparison with limestones. Jurassic lithotypes are characterized by similar development, but differ in
degree of recrystallization.

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the  northern  Boreal  Realm  in  the  NW  and  the  Tethyan
Realm in the SE is suggested by the composition of ammo-
nite  assemblages.  The  marine  ways,  which  led  probably
through the Paris Basin and Germany connecting the Tethyan
Realm with the Boreal Realm (Fig. 4), were first opened dur-
ing the Callovian (Page et al. 2009). During the Late Callov-
ian,  the  Boreal  sea  penetrated  into  Bohemia  from  the  north
via  the  Labe  zone  and  formed  the  Moravian  (Saxonian)
strait, which represents one of the most important migration
routes in Europe during the Late Jurassic (Eliáš 1981).

Systematic paleontology

Order: Ammonitida Zittel, 1885

Superfamily: Perisphinctoidea Steinmann, 1890

Family: Aulacostephanidae Spath, 1924

Genus: Microbiplices Arkell, 1936

Microbiplices sp.

Fig. 5A

M a t e r i a l : NM-N55; from Šternberk quarry near Brtníky.
M e a s u r e m e n t s : NM-N55: D (phg): 28 mm; Wh/D: 0.30;

Wt/D: 0.35; U/D: 0.40.

D e s c r i p t i o n : Carbonate internal mould with preserved

inner whorls. The umbilical part and lateral sides are not pre-
served.  Evolute  type  of  shell  and  regular  rib  bifurcation  is
typical.  Umbilicus  is  wide.  The  whorl  section  is  round  to
oval.  Primaries  have  an  almost  radial  direction.  Secondary
ribs cross the ventral margin fluently without fading.

R e m a r k s : Carbonate internal mould, gently deformed.

Character of ribbing and shell coiling suggests some affinity
to Microbiplices microbiplex figured by Matyja et al. 2006,
fig. 4a and M. microbiplex figured by Schairer 2003 (pl. 2,
fig. 7, 8).

Genus: Prorasenia Schindewolf, 1925

Prorasenia sp.

Fig. 5B—E

M a t e r i a l: NM-N62/1, (fig. 5B); Šternberk near Brtníky.

NM-N183 (fig. 5C); Kyjov near Krásná Lípa. NM-N56
(fig. 5D); Šternberk quarry near Krásná Lípa. NM-N62/2
(fig. 5E); Šternberk quarry near the Brtníky village.

M e a s u r e m e n t s : NM-N62/1 (No. 62): D (max): 22 mm.

NM-N183; D (max): 36 mm. NM-N56; D (max): 27 mm;
Wh/D: 0.23; Wt/D: 0.23, not measured. NM-N62/2; D (max):
12 mm.

D e s c r i p t i o n : Evolute types of shell with low density of

ribbing, only the specimen 5B shows a somewhat higher
density of ribbing. Primaries regularly bifurcate, between
them some ribs trifurcate, which is typical of this genus. It is
visible in the specimens 5C and 5E. Ribs are typically sharp
and they have almost radial direction.

R e m a r k s: NM-N62/1: This specimen suggests a some-

what more prorsiradiate ribs than other figured specimens. It is
similar  to  Rasenia  cf.  hardyi  figured  by  Schweigert  &  Cal-
lomon,  1997,  pl. 7,  fig. 10  and  Rasenia  (Prorasenia)  n. sp.
figured by Geyer, 1961, pl. 1, fig. 8.  NM-N183: Low density
of ribbing suggests the similarity with the Prorasenia sp. fig-

Fig. 4. Map showing paleogeographical situation in Central Europe during the Late Jurassic (modified according to Matyja & Wierzbowski
1995).

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ured  by  Schairer,  2003  (pl. 2,  fig. 12).  Dichotomical  and  tri-
chotomical  branching  of  ribs  is  present.  Both  samples  have
similar  shell  size.  NM-N56:  Specimen  is  deformed  and  in-
complete. It shows some similarity to Prorasenia crenata fig-
ured by Wierzbowski, 2010 (pl. 10, fig. 3, 4). NM-N62/2: One
trichotomically branched rib suggests a similarity with genus
Prorasenia.  The  density  of  ribs  is  not  high.  Prorasenia  cf.
hardyi  figured  by  Schweigert  &  Callomon,  1997  (pl. 7,
fig. 10) has somewhat stronger ribs than the specimen 5E.

Prorasenia cf. bathyschista Koerner, 1963

Fig. 5F—H

M a t e r i a l : NM-N173; (fig. 5F) Kyjov near Krásná Lípa.

CIGP 12; (fig. 5G) former Šternberk quarry. SMGPD CsJ 45
(fig. 5H); Doubice village near Krásná Lípa.

M e a s u r e m e n t s : NM-N173; D (phg-ad): 47 mm; Wh/D:

0.35; Wt/D: 0.32; U/D: 0.45. CIGP 12; D (max): 33 mm;
Wh/D: 0.30; Wt/D: 0.37, not measured. SMGPD CsJ 45;
D (phg): 30 mm; Wh/D: 0.36; Wt/D: 0.37; U/D: 0.42.

D e s c r i p t i o n : Evolute shells with a round whorl cross-

section. Primary ribs are slightly prorsiradiate, secondaries
bend gently backward and form S-shaped deflection.

Re m a r k s : NM-N173: Prorasenia cf. bathyschista closely

resembles the species figured by Matyja & Wierzbowski,
1998 (pl. 2, fig. 4, 5).

Prorasenia cf. crenata Quenstedt, 1887

Fig. 5I

M a t e r i a l : CIGP 10506, from Šternberk quarry.
M e a s u r e m e n t s : D (max): 35 mm; Wh/D: 0.26; Wt/D:

0.28; U/D: 0.37.

D e s c r i p t i o n : Evolute coiling with the umbilical part

and a round whorl cross-section. Primary ribs are gently
prorsiradiate, sharp, and split into two secondaries, which
have backward direction.

R e m a r k s : The specimen is heavily deformed and the

shell is incomplete. The ribbing suggests that it can belong to
Prorasenia. The depicted specimen is of small size (less than
50 mm in diameter), similarly as Prorasenia crenata figured
by Wierzbowski, 2010 (pl. 10, fig. 1—5). The character of rib-
bing is very similar in both specimens. Ribs are weaker then in
P. hardyi figured by Schweigert & Callomon, 1997 (pl. 7,
fig. 10), but the rib shape is similar in both specimens.

O c c u r r e n c e : Prorasenia crenata occurs in the upper

part of the Bimammatum Zone (the Bimammatum and the
Hauffianum Subzones) and the lower part of the Planula Zone.

Genus: Rasenia Salfeld, 1913

Rasenia sp.

Fig. 5J

M a t e r i a l : NM-N179; Kyjov near Krásná Lípa.
M e a s u r e m e n t s : NM-N179; D (phg): 38 mm; Wh/D:

0.26; Wt/D: 0.26; U/D: 0.47.

D e s c r i p t i o n : The character of inner whorls is reduced,

ribbing is not apparent. The whorl cross-section is square-

shaped up to oval with flat lateral sides. Umbilical wall is
very low or absent. Primary ribs have radial direction. On the
very flat ventral side, the secondaries appear with the same
direction as primaries.

R e m a r k s : Gently deformed, internal mould is cracked.

Inner whorl is poorly preserved and does not allow detailed
determination.

Genus: Eurasenia Geyer, 1961

Eurasenia sp.

Fig. 6A—C

M a t e r i a l : NM-N184, (fig. 6A) found in Šternberk quarry

near Krásná Lípa. CIGP 16305, (fig. 6B) found in Hohnstein
quarry (Germany). SMGPD SaJ 7, (fig. 6C) found in Hohnstein
quarry, Germany.

M e a s u r e m e n t s : NM-N184; D (max): 175 mm. CIGP

16305; D (max): 54 mm. SMGPD SaJ 7; D (phg): 28 mm;
Wh/D: 0.31; Wt/D: 0.30; U/D: 0.37.

D e s c r i p t i o n: Very strong, massive and short primary

ribs suggest some affinity to Eurasenia. Our specimens dif-
fer in the direction of ribbing. The specimen 2A has gently
prorsiradiate  ribs,  2B  has  almost  radial  ribs,  and  2C  shows
slightly  rursiradiate  type  of  ribbing.  Umbilicus  is  narrow
with  involute  coiling  of  whorl.  The  whorl  cross-section
seems to be oval to square-shaped.

R e m a r k s : NM-N184: The specimen is similar to Rasenia

(Eurasenia)  pendula  figured  by  Geyer,  1961  (pl. 9,  fig. 7).
R. (E.) frischlini by Geyer, 1961 (pl. 18, fig. 2) is also simi-
lar  to  the  specimen  figured  here.  CIGP  16305:  Rasenia
(Eurasenia)  trimera  figured  by  Geyer,  1961  (pl. 9,  fig. 6)
seems to be very similar to the sample figured here, mainly
by  massive  primary  ribs  and  splitting  into  secondaries.
SMGPD SaJ 7: Very short and massive primary ribs and their
splitting is strikingly similar to Rasenia (Eurasenia) balteata
figured  by  Geyer,  1961  (pl. 18,  fig. 1),  but  differs  by  having
more rursiradiate ribs than the specimen figured here.

Genus: Rasenioides Schindewolf, 1925

Rasenioides sp.

Fig. 6D—H

M a t e r i a l : SMGPD SaJ 12 (fig. 6D), found in Hohnstein

quarry (Germany). SMGPD SaJ 24 (fig. 6E), found in
Hohnstein quarry, (Germany). NM-N96 (fig. 6F), Šternberk
quarry near Brtníky village. SMGPD SaJ 25 (fig. 6G), found
in Hohnstein quarry, (Germany). NM-N60, (fig. 6H); Kyjov
near Krásná Lípa town.

Measurements: SMGPD SaJ 12; D (phg): 29 mm; Wh/D:

0.35;  Wt/D:  0.29;  U/D:  0.42.  SMGPD  SaJ  24;  D (phg):
22 mm,  Wh/D:  0.35;  Wt/D:  0.32;  U/D:  0.28.  NM-N96;
D (phg):  41 mm;  Wh/D:  0.32;  Wt/D:  0.40  U/D:  ?0.45.
SMGPD SaJ 25; D (phg): 30 mm; Wh/D: 0.38; Wt/D: 0.20;
U/D:  0.31.  NM-N60:  D (phg):  40 mm;  Wh/D:  0.39;  Wt/D:
0.28; U/D: 0.33.

D e s c r i p t i o n : Rasenioides is characterized by involute

type of shell with short and strong primary ribs and by high

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density of secondaries. They have less prorsiradiate direc-
tion, sometimes almost radial, than primaries.

R e m a r k s : SMGPD SaJ 12: Rasenioides sp. figured here

resembles  the  Rasenia  (Rasenioides)  lepidula  figured  by
Geyer, 1961 (pl. 8, fig. 5, 6), but the characteristics of prima-
ries are worn away. SMGPD SaJ 24: Rasenia (Rasenioides)
paralepida  (pl. 5,  fig. 10)  and  R.  (R.)  transitoria  (pl. 1,
fig. 6), both figured by Geyer, (1961) differ from this speci-
men figured by higher density of ribbing. A narrow umbili-
cal  part  with  massive  last  whorl  is  typical.  NM-N96:
Massive last whorl with poorly preserved umbilical part sug-
gests the affinity to the Rasenioides. SMGPD SaJ 25: Poorly
preserved  mould  similar  to  Rasenia  (Rasenioides)  transito-
ria figured by Geyer, 1961 (pl. 1, fig. 6). However, the fig-
ured  specimen  differs  by  having  a  more  rursiradiate  rib
direction. NM-N60: Massive outer whorl suggests the affinity
to Rasenioides.

Genus: Vielunia Wierzbowski & Glowniak, 2010

Vielunia sp.

Fig. 6I—L

M a t e r i a l : CIGP 12651: (fig. 6I), found in Šternberk

quarry near Brtníky. NM-N172/1: (fig. 6J), collected on pro-
file in Šternberk quarry near Krásná Lípa. NM-N172/2:
(fig. 6K), collected on profile in Šternberk quarry near Krásná
Lípa. NM-N6: (fig. 6L), found in Šternberk quarry near
Krásná Lípa.

Measurements: CIGP 12651: D (phg): 27 mm; Wh/D:

0.32; Wt/D: 0.35; U/D: 0.33. NM-N172/1: D (phg): 26 mm;
Wh/D: 0.37; Wt/D: 0.34; not measured. NM-N172/2:
D (phg-ad): 25 mm; Wh/D: 0.35; Wt/D: 0.32; not measured.
NM-N6: D (max): 29 mm.

D e s c r i p t i o n : Involute shells with a round or oval whorl

cross-section and a narrow umbilicus. Ribs have radial direc-
tion with a hint of falcoid rib type. Density of ribs is high
and they are thick and blunt.

R e m a r k s : NM-N172/1, 2: Only the last whorl shows

measurable  parameters.  Both  specimens  show  involute
type  of  coiling  with  secondary  filled  umbilical  area.  The
density  of  ribbing  on  the  last  whorl  is  high.  The  figured
specimens  resemble Vielunia dzalosinensis  sp. nov.  figured
by  Wierzbowski  et  al.,  2010  (pl. 9).  NM-N6:  Strongly  de-
formed mould.

Genus: Aulacostephanoides

Aulacostephanoides sp.

Fig. 5K

M a t e r i a l : SMGPD SaJ 3: found in Hohnstein quarry,

Germany.

Measurements: D (max): 38 mm.
D e s c r i p t i o n : A part of massive whorl with high-square

whorl cross-section. Relatively short primary ribs show a
low density. Secondaries have higher density and fade on the
ventral margin, corresponding to the typical character of
Aulacostephanoides.

Genus: Aulacostephanus

Aulacostephanus sp.

Fig. 5L

M a t e r i a l : SMGPD SaJ 15: found in Hohnstein quarry,

(Germany).

Measurements: D (max): 70 mm.
D e s c r i p t i o n : A part of whorl with typically fading ribs

on the ventral margin, with a square-shaped whorl cross-sec-
tion and flattened lateral sides.

R e m a r k s : The weakening of the ribs suggests that this

specimen belongs to Aulacostephanus.

Family: Cardioceratidae Siemiradski, 1891

Genus: Amoeboceras Hyatt, 1900

Amoeboceras ovale Quenstedt, 1849

Fig. 6M

M a t e r i a l : NM-N180: Šternberk quarry.
Measurements: D (phg): 18 mm.
D e s c r i p t i o n : Involute type of shell. The figured speci-

men  with  partly  filled  umbilical  part.  The  whorl  cross-sec-
tion  has  ellipsoid  shape.  The  density  of  ribbing  is  high.  A
keel  is  developed  on  the  ventral  side,  a  typical  character  of
Amoeboceras. Relicts of sutures are recognizable on the lat-
eral side.

R e m a r k s : The specimen figured here shows a similarity

to Amoeboceras ovale figured by Gygi (2000), but the latter
has more flattened lateral sides. This specimen also resem-
bles A. ovale figured by Matyja & Wierzbowski, 1998 (pl. 1,
fig. 5, 6).

O c c u r r e n c e : Amoeboceras ovale occurs in the Regu-

lare and Rosenkrantzi Zones in the Boreal Realm and in the
upper part of the Bifurcatus Zone (Grossouvrei Subzone) up
to the lower part of the Bimammatum Zone (Hypselum Sub-
zone in the Sub-Mediterranean Realm).

Subgenus: Plasmatites Buckman, 1925

Amoeboceras (Plasmatites) praebauhini Salfeld, 1913

Fig. 6N

M a t e r i a l : CIGP 10524: Šternberk quarry.
Measurements: D (phg): 28 mm.
D e s c r i p t i o n : Involute coiling of shell, with secondarily

filled umbilical part. The specimen suggests an inversely
egg-shaped whorl cross-section. A typical keel is partly well
preserved.

R e m a r k s : Gently deformed and cracked carbonate

mould. Some ribs show variable direction caused by defor-
mation. The specimen figured shows a similarity with A.
praebauhini depicted by Atrops et al., 1993 (pl. 1, fig. 15—17).
A. bauhini figured by Schweigert & Callomon, 1997 (pl. 1)
suggests a similar deflection of ribs as the specimen figured
here.

Occurrence: Amoeboceras (Plasmatites) praebauhini oc-

curs in the uppermost part of the Bimammatum Zone (Hauf-

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fianum Subzone) and in the lower part of the Planula Zone in
the Sub-Mediterranean Realm. In the Sub-Boreal Realm,
taxon frequently occurs in the Baylei Zone (Densicostata and
Normandiana Subzones) as well as in the Bauhini and Kit-
chini Zones (Boreal Realm).

Geographical distribution and stratigraphic ranges

of Aulacostephanidae

Aulacostephanid ammonites appeared at the beginning of

the  Late  Oxfordian  and  became  extinct  at  the  end  of  the
Kimmeridgian  (Page  2008).  In  addition  to  Sub-Boreal  re-
gions, Aulacostephanidae were found in some Sub-Mediter-
ranean  regions,  such  as  southern  Germany  (Swabian  Alb,
Franconian  Alb),  Swiss  Jura  and  Central  Poland  (Schneid
1940;  Dabrowska  1984;  Gygi  2000,  2003;  Matyja  &
Wierzbowski  2002;  Glowniak  &  Wierzbowski  2007;  Hor-
nung 2009; Moliner 2009; Glowniak et al. 2010). Species of
this family were recorded also in the Boreal Realm, namely
in East Greenland, Norwegian Sea shelf, Spitsbergen, Franz
Josef Land, western and northern Siberia, British Columbia
and Canadian Arctic Archipelago (Mesezhnikov & Shulgina
1982;  Mesezhnikov  1984,  1988;  Birkelund  &  Callomon
1985; Wierzbowski 1989). However, cardioceratids are domi-
nant  in  the  Boreal  Realm  and  aulacostephanids  are  mostly
restricted to a few stratigraphic intervals (Rogov 2012). Aula-
costephanids  are  rare  or  entirely  missing  at  most  high-lati-
tude  sites,  for  example,  in  northern  Siberia  (Mesezhnikov
1984; Rogov & Wierzbowski 2009; Wierzbowski & Rogov
2013).

Eight aulacostephanid genera (Microbiplices, Prorase-

nia,  Rasenia,  Eurasenia,  Rasenioides,  Vielunia,  Aula-
costephanoides  and  Aulacostephanus)  identified  in  the
Lower  Kimmeridgian  associations  of  northern  Bohemia
represent new aulacostephanid occurrences in the Bohemian
Massif.  The  replacement  of  Microbiplices  by  younger
Eurasenia,  Prorasenia  or  Rasenioides  can  correspond  to
the Oxfordian/Kimmeridgian boundary interval in the Sub-
Boreal Realm (northern France, northern Germany and Great
Britain – Wright 2010). In contrast, the existence of these
aulacostephanid  taxa  indicates  the  Late  Oxfordian  in  the
Sub-Mediterranean  Realm  (German-Polish  branch).  Aula-
costephanids  appeared  in  Subpolar  Urals  during  the  latest
Oxfordian (Mesezhnikov 1984), and during the Early Kim-
meridgian became common in western and northern Siberia
(Rogov & Wierzbowski 2009).

Vielunia, first described by Wierzbowski (2010), occurs in

the upper part of the Bimammatum Zone and continues into
the  Planula  Zone,  corresponding  to  the  lowermost  Kim-
meridgian in the Sub-Boreal and Boreal regions. The genus
Rasenia and related genera (i.e. Prorasenia, Eurasenia, and
Rasenioides) found in the Bohemian Massif belong to Sub-
Boreal ammonites. Rasenia has a relatively broad geographi-
cal  range,  covering  Greenland,  Spitsbergen,  the  Norwegian
Sea,  Franz  Josef  Land,  western  Siberia,  northern  Siberia,
Great  Britain,  France,  Germany,  Poland,  Russian  Platform,
British  Columbia  and  Arctic  Canada  (Tornquist  1896;
Schneid 1940; Birkelund et al. 1978, 1983; Callomon & Bir-

kelund  1980;  Mesezhnikov  1984;  Birkelund  &  Callomon
1985; Matyja et al. 2006; Wierzbowski et al. 2010). There-
fore,  aulacostephanids  from  northern  Bohemia  and  Saxony
thus seem to have a close relationship with ammonites repre-
senting  the  Sub-Mediterranean  branch  frequently  occurring
in southern Germany and Poland.

Aulacostephanoides and Aulacostephanus belong to the

youngest  genera  of  the  family  Aulacostephanidae  described
here. Aulacostephanus was recorded in Sub-Boreal and Bo-
real Realm as well as in some Sub-Mediterranean regions. It
occurs in France, Great Britain, Switzerland, Germany, Nor-
way,  East  Greenland,  Russian  Platform,  western  and  north-
ern  Siberia  (de  Loriol  1874;  Ziegler  1962;  Birkelund  et  al.
1978,  1983;  Mesezhnikov  1984;  Callomon  &  Birkelund
1985).

Geographical distribution and stratigraphic ranges

of Cardioceratidae

The Cardioceratid genus Amoeboceras was described from

the Barents Sea, the Norwegian Sea shelf, Spitsbergen, west-
ern  Siberia,  northern  Siberia,  Russian  Platform,  and  East
Greenland,  (Sykes  &  Callomon  1979;  Birkelund  &  Cal-
lomon 1985; Wierzbowski & Smelror 1993; Wierzbowski et
al.  2002;  Meledina  2006;  Rogov  &  Wierzbowski  2009;
Glowniak  et  al.  2010;  Rogov  2010),  as  well  as  from  Scot-
land,  northern  and  central  Poland,  southern  Germany,  and
Switzerland  (Matyja  &  Wierzbowski  1994,  2002;  Schwei-
gert  &  Callomon  1997;  Gygi  2000;  Matyja  et  al.  2006;
Wierzbowski  et  al.  2010).  The  occurrence  of  Amoeboceras
in northern Bohemia corresponds to the equatorward migra-
tion of Boreal ammonites close to the Oxfordian/Kimmerid-
gian  boundary.  In  the  Polish  Jura  Chain  and  in  southern
Germany,  Amoeboceras  (Plasmatites)  praebauhini  appears
in  the  upper  part  of  the  Sub-Mediterranean  Bimammatum
Zone (Hauffianum Subzone) and continues into the Planula
Zone. This stratigraphic range correlates with the Early Kim-
meridgian Baylei and Bauhini Zones in Sub-Boreal and Bo-
real  regions.  A.  ovale  represents  an  older  species  occurring
during  the  Late  Oxfordian  and  stratigraphically  precedes
Amoeboceras (Plasmatites) praebauhini.

Discussion

Ammonite assemblages sharing many species in common

occur in the Polish Jura Chain, Holy Cross Mountains, Swa-
bian  Alb  and  Franconian  Alb  (Wierzbowski  1978;  Mali-
nowska  1991;  Schweigert  &  Callomon  1997;  Matyja  &
Wierzbowski 2000, 2002; Wierzbowski et al. 2010). Ammo-
nites  occur  in  deep  neritic  sponge  megafacies  in  the  Polish
Jura  Chain  as  well  as  in  southern  Germany,  and  in  more
shallow deposits of the carbonate platform of the Holy Cross
Mts.  The  connection  between  Poland  and  Germany  that
crossed the Bohemian Massif is assumed to have been active
during  the  Late  Jurassic  (Matyja  &  Wierzbowski  1995,
2002; Schairer & Schlampp 2003). The connection between
these two areas enabled a migration of Boreal and Sub-Boreal

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THICA, 2014, 65, 5, 375—386

ammonites  across  the  Bohemian  Massif  southward  to  the
Tethyan  Realm  at  the  Oxfordian/Kimmeridgian  transition
(Atrops  et  al.  1993;  Matyja  &  Wierzbowski  1994,  2000).
Temporal  proximity  and  lithological  similarity  of  the  Late
Jurassic sediments in the North Bohemian and Moravian ar-
eas  (i.e.  Brno  vicinity,  Olomučany  village,  etc.)  evokes  the
possibility of a paleogeographic connection between both re-
gions. However, the direct evidence is still missing because
no Jurassic sediments are preserved between these regions.

Conclusions

The Upper Jurassic marine deposits in northern Bohemian

Massif  are  rarely  preserved  and  limited  to  a  few  small  out-
crops, being mostly covered by Cretaceous and Tertiary de-
posits.  The  stretched  fault  system  associated  with  the
Lusatian  Fault,  reactivated  multiple  times,  does  not  allow  a
discrimination  of  the  original  character  of  shelf  and  basinal
deposits.  However,  ammonites  recovered  from  a  few  out-
crops and preserved in museum collections allow taxonomic
determinations and have implications for paleobiogeographic
pathways  during  the  Late  Jurassic.  Aulacostephanid  ammo-
nites suggest that the Late Jurassic carbonates belong to the
Upper Oxfordian and the Lower Kimmeridgian.

1. The appearance of typical Lower Kimmeridgian Boreal

and Sub-Boreal assemblages in northern Bohemia, repre-
sented by Prorasenia, Rasenioides or Eurasenia, marks the
Oxfordian/Kimmeridgian boundary.

2. With the exception of Prorasenia bathyschista Koerner

figured by Bruder in 1882, ammonites of family Aulaco-
stephanidae are described for the first time from the northern
Bohemian Massif.

3. The presence of Boreal and Sub-Boreal taxa in the

northern Bohemian Massif probably reflects the equatorward
migration of cold-water ammonites close to the Oxfordian/
Kimmeridgian boundary. Taxa occurring in northern Bohe-
mia show affinity to those occurring in the Polish Jura Chain
and southern Germany. This supports the presence of a ma-
rine connection between Poland and Germany across the Bo-
hemian Massif (Matyja & Wierzbowski (1995).

Acknowledgments: This paper was supported by Charles

University  Grant  Agency,  Project  No. 5947/2012,  “Upper
Jurassic  ammonites  of  northern  Bohemia  and  their  signifi-
cance”. The author is indebted to Dr. Jan Sklenář (National
Museum,  Prague)  for  preparing  excellent  conditions  during
my  research  in  the  National  Museum.  I  also  thank  Dr.
Mikhail  Rogov  (GIN  RAS  Moscow),  Dr.  Andrzej
Wierzbowski  (IG,  University  of  Warsaw)  and  Dr.  Günter
Schweigert  (SMN,  Stuttgart)  for  rigorous  help  with  the  de-
termination  of  ammonite  specimens.  Dr.  Markus  Wilmsen
allowed  me  access  to  the  ammonite  collections  of  the  Na-
tional Museum in Dresden, Germany. I would like to thank
the  Institute  of  Geology  and  Paleontology,  Faculty  of  Sci-
ence,  Charles  University  in  Prague.  I  especially  wish  to
thank  Dr.  Martin  Koš ák  for  very  important  and  rewarding
consultation  and  supervision.  Dr.  Martin  Mazuch  and  Dr.
Radek  Vodrážka  helped  with  access  to  fossil  collections.

Finally, I would like to thank Dr. to Adam Tomašových for
valuable tutorial and final improvement of the English and
Mr. Alan Leath for English corrections.

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